The effects of diet-shifting from invertebrates towards fruit on the condition of autumn-migrant Catharus thrushes.

Migration is an energetically challenging and risky life history stage for many animals, but could be supported by dietary choices en route, which may create opportunities to improve body and physiological condition. However, proposed benefits of diet shifts, such as between seasonally available invertebrates and fruits, have received limited investigation in free-living animals. We quantified diet composition and magnitude of autumn diet shifts over two time periods in two closely-related species of migratory songbirds on stopover in the northeastern U.S. (Swainson's thrush [Catharus ustulatus], long-distance migrant, N = 83; hermit thrush [C. guttatus], short-distance migrant, N = 79) and used piecewise structural equation models to evaluate the relationships among (1) migration timing, (2) dietary behavior, and (3) morphometric and physiological condition indices. Tissue isotope composition indicated that both species shifted towards greater fruit consumption. Larger shifts in recent weeks corresponded to higher body condition in Swainson's, but not hermit thrushes, and condition was more heavily influenced by capture date in Swainson's thrushes. Presence of "high-antioxidant" fruits in fecal samples was unrelated to condition in Swainson's thrushes and negatively related to multiple condition indices in hermit thrushes, possibly indicating the value of fruits during migration is related more to their energy and/or macronutrient content than antioxidant content. Our results suggest that increased frugivory during autumn migration can support condition, but those benefits might depend on migration strategy: a longer-distance, more capital-dependent migration strategy could require stricter regulation of body condition aided by increased fruit consumption.

Generalist and topologically central avian frugivores promote plant invasion unequally across land-bridge islands.

Seed dispersal by frugivorous birds facilitates plant invasions, but it is poorly known how invasive plants integrate into native communities in fragmented landscapes. We surveyed plant-frugivore interactions, including an invasive plant (Phytolacca americana), on 22 artificial land-bridge islands (fragmented forests) in the Thousand Island Lake, China. Focusing on frugivory interactions that may lead to seed dispersal, we built ecological networks of studied islands both at the local island (community) and at landscape (metacommunity) levels. On islands with P. americana, we found that P. americana impacted local avian frugivory networks more on islands with species-poor plant communities and on isolated islands. Moreover, as P. americana interacted mainly with local core birds (generalists), this indicates reduced seed dispersal of native plants on invaded islands. At the landscape level, P. americana had established strong interactions with generalist birds that largely maintain seed-dispersal functions across islands, as revealed by their topologically central roles both in the regional plant-bird trophic network and in the spatial metanetwork. This indicates that generalist frugivorous birds may have facilitated the dispersal of P. americana across islands, making P. americana well integrated into the plant-frugivore mutualistic metacommunity. Taken together, our study demonstrates that the impact of plant invasion is context-dependent and that generalist native frugivores with high dispersal potential may accelerate plant invasion in fragmented landscapes. These findings highlight the importance of taking the functional roles of animal mutualists and habitat fragmentation into account when managing plant invasions and their impact on native communities.

Linking pollen limitation and seed dispersal effectiveness.

Seed production and dispersal are crucial ecological processes impacting plant demography, species distributions and community assembly. Plant-animal interactions commonly mediate both seed production and seed dispersal, but current research often examines pollination and seed dispersal separately, which hinders our understanding of how pollination services affect downstream dispersal services. To fill this gap, we propose a conceptual framework exploring how pollen limitation can impact the effectiveness of seed dispersal for endozoochorous and myrmecochorous plant species. We summarize the quantitative and qualitative effects of pollen limitation on plant reproduction and use Optimal Foraging Theory to predict its impact on the foraging behaviour of seed dispersers. In doing so, we offer a new framework that poses numerous hypotheses and empirical tests to investigate links between pollen limitation and seed dispersal effectiveness and, consequently, post-dispersal ecological processes occurring at different levels of biological organization. Finally, considering the importance of pollination and seed dispersal outcomes to plant eco-evolutionary dynamics, we discussed the implications of our framework for future studies exploring the demographic and evolutionary impacts of pollen limitation for animal-dispersed plants.

Rasgos funcionales de frutos con importancia particular para los dispersores de semillas en el bosque seco tropical

Introducción: La frugivoría es un proceso ecológico determinante para la estructuración y regeneración de los bosques. En los trópicos, donde la diversidad de plantas y animales frugívoros es alta, las relaciones interespecíficas son complejas y requieren estudio. Objetivo: Identificar las especies de plantas ecológicamente importantes en dos redes de interacción, y el papel de los rasgos funcionales de los frutos en esas interacciones en un bosque seco. Métodos: Recolectamos 10 frutos por planta de 10 plantas de cada especie de interés en un bosque seco colombiano, calculamos el índice de importancia de las plantas a partir de la relevancia de aves y mamíferos frugívoros en la estructura de las redes. Esta relevancia se relaciona directamente con el potencial del animal como dispersor efectivo de semillas. Utilizamos modelos lineales generalizados para estimar el tamaño, color, estrato, y tipo de pulpa, en el índice. Resultados: Las plantas más importantes son especies de los géneros Miconia, Ficus, Cecropia, Bursera, Casearia y Trichilia, también identificadas como recursos importantes para los frugívoros de los trópicos en otros estudios. Las plantas con frutos carnosos, rojos y de menor tamaño son los mejores para dispersores de semillas. El índice de importancia de las plantas tiene alta variación; esto sugiere que un conjunto de especies frugívoras beneficiadas por cada especie de planta tiene una contribución diferenciada en procesos ecológicos derivados de la dispersión de semillas. Conclusiones: Programas de restauración para este tipo de bosque tropical seco debería incluir una variedad de plantas, incluyendo especies con frutos pequeños, rojos y carnosos.

Introduction: Frugivory is a pivotal ecological process for the structure and regeneration of forests. In the tropics, where the diversity of plants and frugivorous animals is high, interspecific relationships in the interaction networks are complex and need study. Objective: To identify ecologically important plant species in two interaction networks, and the role of functional fruit traits in those interactions in a dry forest. Methods: We collected 10 fruits per plant from 10 plants of each species of interest in a Colombian dry forest, we calculated the Plant Importance Index based on the bird and mammal frugivores relevance for network structure. This relevance is directly related to the animal's potential as effective seed dispersers. We used generalized linear models to estimate the effect of fruit size, color, stratum, and type of pulp, on the index. Results: The most important plants are species of the genera Miconia, Ficus, Cecropia, Bursera, Casearia and Trichilia, also identified as important resources for tropical frugivores in other studies. Plants with small, red, and fleshy fruits are the best for seed dispersers. The plant importance index has a high variation; this suggests that the set of frugivore species benefited by each plant species has a differential contribution to the ecological processes derived from seed dispersal. Conclusions: Restoration programs for this kind of tropical dry forest should include a variety of plants, including species with small, red, and fleshy fruits.

What drives seed dispersal effectiveness?

Seed dispersal is a critical phase in plant reproduction and forest regeneration. In many systems, the vast majority of woody species rely on seed dispersal by fruit-eating animals. Animals differ in their size, movement patterns, seed handling, gut physiology, and many other factors that affect the number of seeds they disperse, the quality of treatment each individual seed receives, and consequently their relative contribution to plant fitness. The seed dispersal effectiveness framework (SDE) was developed to allow systematic and standardized quantification of these processes, offering a potential for understanding the large-scale dynamics of animal-plant interactions and the ecological and evolutionary consequences of animal behavior for plant reproductive success. Yet, despite its wide acceptance, the SDE framework has primarily been employed descriptively, almost always in the context of local systems. As such, the drivers of variation in SDE across systems and the relationship between its components remain unknown. We systematically searched studies that quantified endozoochorous SDE for multiple animal species dispersing one or more plant species in a given system and offered an integrative examination of the factors driving variation in SDE. Specifically, we addressed three main questions: (a) Is there a tradeoff between high dispersal quality and quantity? (b) Does animal body mass affect SDE or its main components? and (c) What drives more variation in SDE, seed dispersal quality, or quantity? We found that: (a) the relationship between quality and quantity is mediated by body size; (b) this is the result of differential relationships between body mass and the two components, while total SDE is unaffected by body mass; (c)neither quality nor quantity explain more variance in SDE globally. Our results also highlight the need for more standardized data to assess large-scale patterns in SDE.

Fruit and seed traits and vertebrate-fruit interactions of tree species occurring in Guyana, Suriname, and French Guiana.

Seed dispersal is widely considered an important mechanism for the conservation of plant diversity. In tropical regions, over 80% of woody plant species are dispersed by vertebrates, often through the consumption of fruits. Our understanding of what drives interactions between vertebrates and fruits is limited. Through a systematic literature search, we compiled a database of fruit and seed traits and vertebrate-fruit interactions for tree and vertebrate species occurring in the Guianas, with the aim of facilitating research into seed dispersal and seed predation of tree species in the Guianas. The database was compiled by extracting data from 264 published sources. It consists of 21,082 records, of which 19,039 records contain information about 19 different fruit and seed traits belonging to 1622 different tree species. The other 2043 records contain information on vertebrate-fruit interactions between 161 vertebrate species and 464 tree species. Our analyses showed a taxonomic bias, particularly in the interaction data, toward large-bodied vertebrates, with most interactions recorded for the bearded saki (Chiropotes chiropotes), followed by the lowland tapir (Tapirus terrestris). For plants we found an overrepresentation of the Sapotaceae and Moraceae families and an underrepresentation of the Rubiaceae, Myrtaceae, and Lauraceae families in the interactions. There are no copyright restrictions on the data set; please cite this publication when using these data.

Seed dispersal syndrome predicts ethanol concentration of fruits in a tropical dry forest.

Studying fruit traits and their interactions with seed dispersers can improve how we interpret patterns of biodiversity, ecosystem function and evolution. Mounting evidence suggests that fruit ethanol is common and variable, and may exert selective pressures on seed dispersers. To test this, we comprehensively assess fruit ethanol content in a wild ecosystem and explore sources of variation. We hypothesize that both phylogeny and seed dispersal syndrome explain variation in ethanol levels, and we predict that fruits with mammalian dispersal traits will contain higher levels of ethanol than those with bird dispersal traits. We measured ripe fruit ethanol content in species with mammal- (n = 16), bird- (n = 14) or mixed-dispersal (n = 7) syndromes in a Costa Rican tropical dry forest. Seventy-eight per cent of fruit species yielded measurable ethanol concentrations. We detected a phylogenetic signal in maximum ethanol levels (Pagel's λ = 0.82). Controlling for phylogeny, we observed greater ethanol concentrations in mammal-dispersed fruits, indicating that dispersal syndrome helps explain variation in ethanol content, and that mammals may be more exposed to ethanol in their diets than birds. Our findings further our understanding of wild fruit ethanol and its potential role as a selective pressure on frugivore sensory systems and metabolism.

Take me for a ride: Herbivores can facilitate plant reinvasions.

Herbivores shape plant invasions through impacts on demography and dispersal, yet only demographic mechanisms are well understood. Although herbivores negatively impact demography by definition, they can affect dispersal either negatively (e.g., seed consumption), or positively (e.g., caching). Exploring the nuances of how herbivores influence spatial spread will improve the forecasting of plant movement on the landscape. Here, we aim to understand how herbivores impact how fast plant populations spread through varying impacts on plant demography and dispersal. We strive to determine whether, and under what conditions, we see net positive effects of herbivores, in order to find scenarios where herbivores can help to promote spread. We draw on classic invasion theory to develop a stage-structured integrodifference equation model that incorporates herbivore impacts on plant demography and dispersal. We simulate seven herbivore "syndromes" (combinations of demographic and/or dispersal effects) drawn from the literature to understand how increasing herbivore pressure alters plant spreading speed. We find that herbivores with solely negative effects on plant demography or dispersal always slow plant spreading speed, and that the speed slows monotonically as herbivore pressure increases. However, we also find that plant spreading speed can be hump shaped with respect to herbivore pressure: plants spread faster in the presence of herbivores (for low herbivore pressure) and then slower (for high herbivore pressure). This result is robust, occurring across all syndromes in which herbivores have a positive effect on plant dispersal, and is a sign that the positive effects of herbivores on dispersal can outweigh their negative effects on demography. For all syndromes we find that sufficiently high herbivore pressure results in population collapse. Thus, our findings show that herbivores can speed up or slow down plant spread. These insights allow for a greater understanding of how to slow invasions, facilitate native species recolonization, and shape range shifts with global change.

Africa as an evolutionary arena for large fruits.

Strong paleoclimatic change and few Late Quaternary megafauna extinctions make mainland Africa unique among continents. Here, we hypothesize that, compared with elsewhere, these conditions created the ecological opportunity for the macroevolution and geographic distribution of large fruits. We assembled global phylogenetic, distribution and fruit size data for palms (Arecaceae), a pantropical, vertebrate-dispersed family with > 2600 species, and integrated these with data on extinction-driven body size reduction in mammalian frugivore assemblages since the Late Quaternary. We applied evolutionary trait, linear and null models to identify the selective pressures that have shaped fruit sizes. We show that African palm lineages have evolved towards larger fruit sizes and exhibited faster trait evolutionary rates than lineages elsewhere. Furthermore, the global distribution of the largest palm fruits across species assemblages was explained by occurrence in Africa, especially under low canopies, and extant megafauna, but not by mammalian downsizing. These patterns strongly deviated from expectations under a null model of stochastic (Brownian motion) evolution. Our results suggest that Africa provided a distinct evolutionary arena for palm fruit size evolution. We argue that megafaunal abundance and the expansion of savanna habitat since the Miocene provided selective advantages for the persistence of African plants with large fruits.

Feeding ecology of monk sakis (Pithecia monachus) in a seasonally flooded forest in western Amazonia.

Sakis (genus Pithecia) are frugivorous primates with a preference for seeds that complete their diet with leaves and insects. Fruit pulp and seeds are known to have different nutritional characteristics that change during the process of ripening. The consumption of seeds can be an adaptation to changes in resource availability, as unripe seeds are a more steadily available resource than ripe pulp or young leaves. Here, we present the first study of the feeding ecology of monk sakis (Pithecia monachus). We investigated dietary composition and identified important feeding plants in a seasonally flooded forest within the Área de Conservación Regional Comunal Tamshiyacu-Tahuayo in Peruvian Amazonia. Throughout 20 months, we followed groups of monk sakis by foot and canoe and recorded 459 feeding events. Seeds were the most frequently consumed food item (49%), followed by pulp (mesocarp, pericarp or aril; 25%) and arthropods (22%). Leaves, bark, and flowers were ingested only sporadically. The importance of ripe seeds and arthropods in the diet of the monk sakis differed from other studies: we recorded the consumption of mostly ripe seeds and the share of arthropods was relatively high.

When lizards shift to a more plant-based lifestyle: The macroevolution of mutualistic lizard-plant-interactions (Squamata: Sauria/Lacertilia).

Pollination and seed dispersal of plants by animals are key mutualistic processes for the conservation of plant diversity and ecosystem functioning. Although different animals frequently act as pollinators or seed dispersers, some species can provide both functions, so-called 'double mutualists', suggesting that the evolution of pollination and seed dispersal may be linked. Here, we assess the macroevolution of mutualistic behaviours in lizards (Lacertilia) by applying comparative methods to a phylogeny comprising 2,838 species. We found that both flower visitation (potential pollination; recorded in 64 species [2.3% of total] across 9 families) and seed dispersal (recorded in 382 species [13,5% of total] across 26 families) have evolved repeatedly in Lacertilia. Furthermore, we found that seed dispersal activity pre-dated flower visitation and that the evolution of seed dispersal activity and flower visitation was correlated, illustrating a potential evolutionary mechanism behind the emergence of double mutualisms. Finally, we provide evidence that lineages with flower visitation or seed dispersal activity have higher diversification rates than lineages lacking these behaviours. Our study illustrates the repeated innovation of (double) mutualisms across Lacertilia and we argue that island settings may provide the ecological conditions under which (double) mutualisms persist over macroevolutionary timescales.

Urbanization shapes phenotypic selection of fruit traits in a seed-dispersal mutualism.

Urbanization is currently one of the trademarks of the Anthropocene, accelerating evolutionary processes and reshaping ecological interactions over short time scales. Species interactions represent a fundamental pillar of diversity that is being altered globally by anthropogenic change. Urban environments, despite their potential impact, have seldom been studied in relation to how they shape natural selection of phenotypic traits in multispecies interactions. Using a seed-dispersal mutualism as a study system, we estimated the regime and magnitude of phenotypic selection exerted by frugivores on fruit and seed traits across three plant populations with different degrees of urbanization (urban, semiurban, and rural). Urbanization weakened phenotypic selection via an indirect positive impact on fruit production and fitness and, to a lesser extent, through a direct positive effect on species visitation rates. Our results show that urban ecosystems may affect multifarious selection of traits in the short term and highlight the role of humans in shaping eco-evolutionary dynamics of multispecies interactions.

The turnover of plant-frugivore interactions along plant range expansion: consequences for natural colonization processes.

Plant-animal mutualisms such as seed dispersal are key interactions for sustaining plant range shifts. It remains elusive whether the organization of interactions with seed dispersers is reconfigured along the expansion landscape template and, if so, whether its effects accelerate or slow colonization. Here we analyse plant-frugivore interactions in a scenario of rapid population expansion of a Mediterranean juniper. We combined network analyses with field surveys, sampling interactions between individual plants and frugivores by DNA-barcoding and phototrapping over two seasons. We assess the role of intrinsic and extrinsic intraspecific variability in shaping interactions and we estimate the individual plant contributions to the seed rain. The whole interaction network was highly structured, with a distinct set of modules including individual plants and frugivore species arranged concordantly along the expansion gradient. The modular configuration was partially shaped by individual neighbourhood context (density and fecundity) and phenotypic traits (cone size). Interaction reconfiguration resulted in a higher and more uneven propagule contribution, with most effective dispersers having a prominent role at the colonization front stand, where a distinct subset of early arriving plants dominated the seed rain. Our study offers new insights into the key role of mutualistic interactions in colonization scenarios by promoting fast plant expansion processes.

Invasive Plant Species Driving the Biotic Homogenization of Plant-Frugivore Interactions in the Atlantic Forest Biodiversity Hotspot.

Although biological invasions are a common and intensively studied phenomenon, most studies often ignore the biotic interactions that invasive species play in the environment. Here, we evaluated how and why invasive plant species are interconnected within the overall frugivory network of the Brazilian Atlantic Forest, an important global biodiversity hotspot. To do this, we used the recently published Atlantic Frugivory Dataset to build a meta-network (i.e., a general network made of several local networks) that included interactions between 703 native and invasive plant species and 331 frugivore species. Using tools derived from complex network theory and a bootstrap simulation approach, we found that the general structure of the Atlantic Forest frugivory network (i.e., nestedness and modularity) is robust against the entry of invasive plant species. However, we observed that invasive plant species are highly integrated within the frugivory networks, since both native and invasive plant species play similar structural roles (i.e., plant status is not strong enough to explain the interactive roles of plant species). Moreover, we found that plants with smaller fruits and with greater lipid content play a greater interactive role, regardless of their native or invasive status. Our findings highlight the biotic homogenization involving plant-frugivore interactions in the Atlantic Forest and that the impacts and consequences of invasive plant species on native fauna can be anticipated based on the characteristics of their fruits.

Multiscale phenological niches of seed fall in diverse Amazonian plant communities.

Phenology has long been hypothesized as an avenue for niche partitioning or interspecific facilitation, both promoting species coexistence. Tropical plant communities exhibit striking diversity in reproductive phenology, but many are also noted for large synchronous reproductive events. Here we study whether the phenology of seed fall in such communities is nonrandom, the temporal scales of phenological patterns, and ecological factors that drive reproductive phenology. We applied multivariate wavelet analysis to test for phenological synchrony versus compensatory dynamics (i.e., antisynchronous patterns where one species' decline is compensated by the rise of another) among species and across temporal scales. We used data from long-term seed rain monitoring of hyperdiverse plant communities in the western Amazon. We found significant synchronous whole-community phenology at multiple timescales, consistent with shared environmental responses or positive interactions among species. We also observed both compensatory and synchronous phenology within groups of species (confamilials) likely to share traits and seed dispersal mechanisms. Wind-dispersed species exhibited significant synchrony at ~6-month scales, suggesting these species might share phenological niches to match the seasonality of wind. Our results suggest that community phenology is shaped by shared environmental responses but that the diversity of tropical plant phenology may partly result from temporal niche partitioning. The scale-specificity and time-localized nature of community phenology patterns highlights the importance of multiple and shifting drivers of phenology.

Bird preferences for fruit size, but not color, vary in accordance with fruit traits along a tropical elevational gradient.

Birds constitute one of the most important seed dispersal agents globally, especially in the tropics. The feeding preferences of frugivorous birds are, therefore, potentially of great ecological importance. A number of laboratory-based and observational studies have attempted to ascertain the preferences of certain bird species for certain fruit traits. However, little attention has been paid to community-wide preferences of frugivorous birds and the impact this may have on fruit traits on a broader scale. Here, we used artificial fruits of different colors and sizes to investigate community-wide fruit trait preferences of birds at three sites along an elevational gradient in Papua New Guinea. We recorded attack rates on artificial fruits as visible impressions made by a bird's beak during a feeding attempt. We also measured the colors and sizes of real fruits at each site, and the gape widths of frugivorous birds, allowing for comparisons between bird feeding preferences and bird and fruit traits. Regardless of elevation, red and purple fruits were universally preferred to green and attacked at similar rates to one another, despite strong elevational patterns in real fruit color. However, elevation had a significant effect on fruit size preferences. A weak, non-significant preference for large fruits was recorded at 700 m, while medium fruits were strongly preferred at 1700 m and small fruits at 2700 m. These patterns mirror those of both real fruit size and frugivorous bird gape width along the gradient, suggesting the potential for selective pressure of birds on fruit size at different elevations.

Ecological Impacts of Exotic Species on Native Seed Dispersal Systems: A Systematic Review.

Exotic species are one of the main threats to biodiversity, leading to alterations in the structure and functioning of natural ecosystems. However, they can sometimes also provide ecological services, such as seed dispersal. Therefore, we assessed the ecological impacts of exotic species on native dispersal systems and the mechanisms underlying the disruption of mutualistic plant-disperser interactions. Exotic species negatively affect dispersal mutualisms by (i) altering dispersal behavior and visitation rates of native dispersers, (ii) predating native dispersers, (iii) transmitting forest pathogens, and (iv) predating seeds. Conversely, positive impacts include the dispersal of native plants, forest regeneration, and native habitat restoration via (i) increasing the visitation rates of frugivorous birds, (ii) facilitating the colonization and establishment of native forest trees, (iii) enhancing forest species seedling survival, and (iv) facilitating seed rain and seedling recruitment of early and late successional native plants. The reviewed studies provide similar results in some cases and opposite results in others, even within the same taxa. In almost all cases, exotic species cause negative impacts, although sometimes they are necessary to ensure native species' persistence. Therefore, exotic species management requires a comprehensive understanding of their ecological roles, since the resulting effects rely on the complexity of native-exotic species interactions.

Life history and chemical defense interact to drive patterns of adaptation in an annual monkeyflower.

Although chemical defenses and herbivore pressure are widely established as key targets and agents of selection, their roles in local adaptation and determining potential evolutionary responses to changing climates are often neglected. Here, we explore fitness differences between 11 rangewide M. guttatus populations in a field common garden experiment and assess the agents and targets of selection driving relative fitness patterns. We use piecewise structural equation models to disentangle associations between chemical defenses, (phenylpropanoid glycosides; PPGs), and life history traits with herbivory and fitness. While the historical environment of populations is not predictive of fitness differences between populations, >90% of variation in fitness can be predicted by the flowering time and foliar PPG defense arsenal of a population. Piecewise structural equation models indicate that life history traits, particularly earlier flowering time, are strongly and directly linked to fitness. However, herbivory, particularly fruit predation, is also an important agent of selection that creates indirect links between fitness and both chemical defenses and life history traits. Our results emphasize the multivariate nature of the agents and targets of selections in producing adaptation and suggest that future responses to selection must navigate a complex fitness landscape.

On the adequacy of fruit removal as a proxy for fitness in studies of bird-mediated phenotypic selection.

PREMISE: In fleshy-fruited plants, fruit removal is widely used as a proxy for plant reproductive success. Nevertheless, this proxy may not accurately reflect the number of seeds dispersed, an assumed better proxy for total fitness (fruit removal × mean number of seeds dispersed per fruit). METHODS: We examined under what circumstances fruit removal can be reliable as a proxy for total fitness when assessing bird-mediated selection on fruit traits. In three populations of the Blue Passionflower (Passiflora caerulea), we used the number of fruits pecked per plant as a surrogate for fruit removal to estimate phenotypic selection on fruit and seed traits, and simulations of the effect of the fruit-seed number trade-off on the number of fruits removed. RESULTS: Fruit removal was a good indicator of fitness, accounting for 55 to 68% of the variability in total fitness, measured as total number of seeds removed. Moreover, multivariate selection analyses on fruit crop size, mean fruit diameter and mean seed number using fruit removal as a fitness proxy yielded similar selection regimes to those using total fitness. Simulations showed that producing more fruits, a lower number of seeds per fruit, and a higher variability in seed number can result in a negative relationship between fruit removal and total fitness. CONCLUSIONS: Our results suggest that fruit removal can be reliably used as a proxy for total fitness when (1) there is a weak fruit number-seed number trade-off, (2) fruit crop size and fruit removal correlate positively, and (3) seed number variability does not largely exceed fruit number variability.

Reciprocity and interaction effectiveness in generalised mutualisms among free-living species.

Mutualistic interactions among free-living species generally involve low-frequency interactions and highly asymmetric dependence among partners, yet our understanding of factors behind their emergence is still limited. Using individual-based interactions of a super-generalist fleshy-fruited plant with its frugivore assemblage, we estimated the Resource Provisioning Effectiveness (RPE) and Seed Dispersal Effectiveness (SDE) to assess the balance in the exchange of resources. Plants were highly dependent on a few frugivore species, while frugivores interacted with most individual plants, resulting in strong asymmetries of mutual dependence. Interaction effectiveness was mainly driven by interaction frequency. Despite highly asymmetric dependences, the strong reliance on quantity of fruit consumed determined high reciprocity in rewards between partners (i.e. higher energy provided by the plant, more seedlings recruited), which was not obscured by minor variations in the quality of animal or plant service. We anticipate reciprocity will emerge in low-intimacy mutualisms where the mutualistic outcome largely relies upon interaction frequency.